中国内脐蠕孢属（Drechslera）及突脐蠕孢属（Exserohilum）真菌分类研究
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摘要
从全国17省市实地采集的3000多份植物标本进行保湿培养或组织分离
法分离中，得到属于两属真菌的菌株200份，经鉴定，得到一个新种一燕
麦生内脐蠕孢Drechslera avenicola B.D.Sun Sun et T．Y Zhang，5个中国新
记录种：安德森内脐蠕孢D.andersenii Lam、黑麦草内脐蠕孢D.phlei
(Graham)Shoem．、席氏内脐蠕孢D.sivanesanii C.Manoharachary et V.R
T．Reddy、穆子突脐蠕孢E frumenracef(Mitra)Leonard & Suggs、小柄突
脐蠕孢E.pedicellatium(Henry)Leonard & Suggs，国内已记录种11个：
燕麦内脐蠕孢D.avenacea(M．A．Curtis ex Cooke)Shoem．、网斑内脐蠕孢
D.dictyoides(Drechsler) Shoem、禾内脐蠕孢D.graminea(Rabenh．ex
Schlecht)Shoem．、大麦网斑内脐蠕孢D.teres(Sacc．)Shoem．、梭形突脐
蠕孢E.fusiforme Alcom、环形突脐蠕孢E.holmii(Luttr．)I．enoard&suggs、
长孢突脐蠕孢E.10ngisporium G．Y．Sun，R．Zhang，M．Q．Zhu&T．’~．Zhang、
尖角突脐蠕孢E.monoceras(Drechslerl)Leonard & Suggs、 芦苇突脐蠕孢
E. phragmatis W．P Wu、嘴突突脐蠕孢E.rostratum(Drechler)Leonard
&suggs、大斑病突脐蠕孢E turcicum(PaSS)Leonard & Suggs。
对种内分生孢子形态变化较大的嘴突突脐蠕孢Erostratum的26个菌株
作了RAPD分析，结果表明，该种种内遗传变异较大，但聚类结果的分组
并不同于形态上的分组，且具有与形态学相似的无序性，分生孢子形态上
的变化不能成为菌株间亲缘关系的标准，应该将该种作为一个群体对待，
分生孢子形态在自然寄主上有些可能较规则，但从整个类群整体来看，在
复杂多变地自然生态条件下，分生孢子形态变异的幅度大，应该以适宜产
孢的一致培养条件下的特征作为分种的主要依据。
本研究的主要成果为，在前人工作基础上，通过系统深入的研究，订正
了两属及其种的拉、汉名称，发现了一个新种，数个新记录种，增加了一
些种的寄主及地理分布及存在状态，使我们能更全面地了解两属真菌在中
国的分布。在研究过程中，始终注意分类方法及种级分类标准的探索、验
证。对培养特征、内脐蠕孢属的产孢条件、分生孢子隔膜发育、萌发等特
性进行了观察，总结出相对稳定、易行的属间及属内种间分类标准如下：
1．属级分类标准
(1)脐部特征是属的主要划分依据，Drechs．1era属真菌的脐点为腔孔型
(atrium)，在观察中可以看到脐点决不突出分生孢子基细胞的光滑外形，有
时甚至可以从侧面看到褐色的脐点凹陷在基细胞内，与孢子基部细胞壁之
间分离，一般较大。
Exseroh，]um属真菌的脐点明显突出，电子显微镜下可见有一特殊的封
套结构(enveloping structure)，此结构在光学显微镜下多数不能观察到，
但在E.monoceras的有些分生孢子可以看到脐点顶端稍有环状加厚。
(2)基部细胞的萌发特征可以很容易的区分Drechslez．a属真菌与
Bipolaris属真菌，但后者与Exserohilum属真菌在分生孢子萌发特性之
间却没有明显区别。
(3)分生孢子颜色对分属有参考意义。
(4)分生孢子梗的形态及产孢方式在区分Drechslera、Bipolaris和
Exserohilume时，光学显微镜下可以区分的特征较少，在实际分类工作中
价值较小。
(5)分生孢子隔膜是假隔膜还是与外壁相连的真隔膜，在区分Exserohilum
与脐点突出的弯孢属Curvularia的种类时，以前是很重要的标准，但此标
准近来受到怀疑。
2．种级分类标准
(1)分生孢子形态大小、隔膜数量、细胞形状大小、次生分生孢子的情况
在分种时有重要意义。一些参数，例如：平均细胞长度、顶部隔膜宽度与
基部隔宽度比等，在个别形态特别相近的种相互区分时有特殊的价值。
(2)分生孢子梗难以单独作为种间区分标准，其长度的变幅常相互交错，
只能作为分种的次要标准。
(3)寄主和培养性状在．Drechslera属的种间区分中较为重要，而在
Exserohilum属内种间无明显的差异。
在我国丰富的自然生态条件下，深入研究肯定还会有该类群真菌的不断
发现。而且，对于嘴突突脐蠕孢种内分生孢子形态的变异的深层原因、内
脐蠕孢属的部分菌株的产孢条件都有待于继续研究。
This study is about the taxonomic research of Drechslera Ito and
Exsehilum Leonard & Suggs in China. About 200 isolates were received
from more than 3000 herbarium specimens, which we have collected from
many provinces of China.
From the isolates, we have found one new species, D. a venicola B. D.
Sun et T. Y. Zhang, five Chinese new record species, D. andersenii Lam,
D. sivanesanii C. Manoharachary et V. R. T. Reddy, E. frumentaeei (Mitra)
Leonard & Suggs, E. pedicellatium (Henry) Leonard &Suggs and eleven
species which had been reported in China, they are , D. avenacea
(M.A.Curtis ex Cooke) Shoem., O. dictyoides (Drechsler) Shoem.,
D. graminea (Rabenh. ex Schlecht.) Shoem., D. phlei (Graham) Shoem.,.
D. teres (Sate.) Shoem., E. fusiforme Alcom, E. holmii (Luttr.) Lenoard
& suggs, E. ]ongisporium G.Y.Sun, R.Zhang, M.Q.Zhu & T.Y.Zhang,
E. monoceras(Drechsler) Leonard & Suggs, E. phragmatis W.P Wu,
E. rostratum (Drechler) Leonard & Suggs and E. turcicum (Pass.) Leonard
& Suggs.
To investigate the genetic variation in E.rostratum, a total of 26
E.rostratum isolates and 7 isolates of other species were analyzed with 10
random amplified polymorphic DNA (RAPD) primers. The results showed
that there was a similar disorderliness with conidia morphology showed. There
was a large variation among all E.rostratum isolates, and character of conidia
was not a critical criteria of relationship between isolates. We should treat all
isolates of E.rostratum as one group since there were no constant crt. The conidia from culture medium shows less variety when
treated all isolates of E.rostratum as a whole, and should be treated as main
criteria of classification of this species.
After many works have been done about this two genera, the main results
of this research are the revisions of the Latin and Chinese name of the genera
and the species in them. We also have found a new species, several new
Chinese record species and broadened the hosts' records and distribution of
some species. All the works will contribute to the further study of this group of
fungi.
In our research work, we ve
payed a constant attention to the taxonomic
methods and the testifying and exploring of criteria to differentiate species. We
studied the culture character of all isolates, the conditions to spur Drechslera
spp to produce conidia on culture medium, the formation of conidia septum,
the germ character of conidia etc, and have concluded a much easier and
feasible criteria for the classification this group of fungi.
In differentiating different genera, the character of hilum plays an
important role. The hilum of Drechslera spp is atrium type hilum, which is
larger and we can see them never protruding from the contour of basal cell
when under optical microscope. Some time we can even see them were in the
basal cell of conidia, and clear of the cell wall. The hilum of Exserohilum spp.
were protruding and had an enveloping structure. But this structure can not be
found using optical microscope. Only that you may found an ring thickness at
the top of the hilum of E.monoceras and several other species. The
germination character of the basal cell of conidia is useful in distinguishing
Drechslera spp. from Bipolaris spp and Exserohilum spp But this character has
little difference between the later two genera. The color of conidia only has an
assitant role in genus classification. The Exserohilum spp have distoseptates
and the Curivularia spp with protruding hilum have euseptates has been seen
as the main difference between them. This criteria is used even it has been
doubted because there is no other criteria better.
In treating with the species, the morphology of conidia is main criteria,
especialy the shape and the size of conidia, the shape of conidia cell, the
conidia septum and the existence of secondary conidiophore and secondary
conidia. Some parameter such as the ratio of width of a
法分离中，得到属于两属真菌的菌株200份，经鉴定，得到一个新种一燕
麦生内脐蠕孢Drechslera avenicola B.D.Sun Sun et T．Y Zhang，5个中国新
记录种：安德森内脐蠕孢D.andersenii Lam、黑麦草内脐蠕孢D.phlei
(Graham)Shoem．、席氏内脐蠕孢D.sivanesanii C.Manoharachary et V.R
T．Reddy、穆子突脐蠕孢E frumenracef(Mitra)Leonard & Suggs、小柄突
脐蠕孢E.pedicellatium(Henry)Leonard & Suggs，国内已记录种11个：
燕麦内脐蠕孢D.avenacea(M．A．Curtis ex Cooke)Shoem．、网斑内脐蠕孢
D.dictyoides(Drechsler) Shoem、禾内脐蠕孢D.graminea(Rabenh．ex
Schlecht)Shoem．、大麦网斑内脐蠕孢D.teres(Sacc．)Shoem．、梭形突脐
蠕孢E.fusiforme Alcom、环形突脐蠕孢E.holmii(Luttr．)I．enoard&suggs、
长孢突脐蠕孢E.10ngisporium G．Y．Sun，R．Zhang，M．Q．Zhu&T．’~．Zhang、
尖角突脐蠕孢E.monoceras(Drechslerl)Leonard & Suggs、 芦苇突脐蠕孢
E. phragmatis W．P Wu、嘴突突脐蠕孢E.rostratum(Drechler)Leonard
&suggs、大斑病突脐蠕孢E turcicum(PaSS)Leonard & Suggs。
对种内分生孢子形态变化较大的嘴突突脐蠕孢Erostratum的26个菌株
作了RAPD分析，结果表明，该种种内遗传变异较大，但聚类结果的分组
并不同于形态上的分组，且具有与形态学相似的无序性，分生孢子形态上
的变化不能成为菌株间亲缘关系的标准，应该将该种作为一个群体对待，
分生孢子形态在自然寄主上有些可能较规则，但从整个类群整体来看，在
复杂多变地自然生态条件下，分生孢子形态变异的幅度大，应该以适宜产
孢的一致培养条件下的特征作为分种的主要依据。
本研究的主要成果为，在前人工作基础上，通过系统深入的研究，订正
了两属及其种的拉、汉名称，发现了一个新种，数个新记录种，增加了一
些种的寄主及地理分布及存在状态，使我们能更全面地了解两属真菌在中
国的分布。在研究过程中，始终注意分类方法及种级分类标准的探索、验
证。对培养特征、内脐蠕孢属的产孢条件、分生孢子隔膜发育、萌发等特
性进行了观察，总结出相对稳定、易行的属间及属内种间分类标准如下：
1．属级分类标准
(1)脐部特征是属的主要划分依据，Drechs．1era属真菌的脐点为腔孔型
(atrium)，在观察中可以看到脐点决不突出分生孢子基细胞的光滑外形，有
时甚至可以从侧面看到褐色的脐点凹陷在基细胞内，与孢子基部细胞壁之
间分离，一般较大。
Exseroh，]um属真菌的脐点明显突出，电子显微镜下可见有一特殊的封
套结构(enveloping structure)，此结构在光学显微镜下多数不能观察到，
但在E.monoceras的有些分生孢子可以看到脐点顶端稍有环状加厚。
(2)基部细胞的萌发特征可以很容易的区分Drechslez．a属真菌与
Bipolaris属真菌，但后者与Exserohilum属真菌在分生孢子萌发特性之
间却没有明显区别。
(3)分生孢子颜色对分属有参考意义。
(4)分生孢子梗的形态及产孢方式在区分Drechslera、Bipolaris和
Exserohilume时，光学显微镜下可以区分的特征较少，在实际分类工作中
价值较小。
(5)分生孢子隔膜是假隔膜还是与外壁相连的真隔膜，在区分Exserohilum
与脐点突出的弯孢属Curvularia的种类时，以前是很重要的标准，但此标
准近来受到怀疑。
2．种级分类标准
(1)分生孢子形态大小、隔膜数量、细胞形状大小、次生分生孢子的情况
在分种时有重要意义。一些参数，例如：平均细胞长度、顶部隔膜宽度与
基部隔宽度比等，在个别形态特别相近的种相互区分时有特殊的价值。
(2)分生孢子梗难以单独作为种间区分标准，其长度的变幅常相互交错，
只能作为分种的次要标准。
(3)寄主和培养性状在．Drechslera属的种间区分中较为重要，而在
Exserohilum属内种间无明显的差异。
在我国丰富的自然生态条件下，深入研究肯定还会有该类群真菌的不断
发现。而且，对于嘴突突脐蠕孢种内分生孢子形态的变异的深层原因、内
脐蠕孢属的部分菌株的产孢条件都有待于继续研究。
This study is about the taxonomic research of Drechslera Ito and
Exsehilum Leonard & Suggs in China. About 200 isolates were received
from more than 3000 herbarium specimens, which we have collected from
many provinces of China.
From the isolates, we have found one new species, D. a venicola B. D.
Sun et T. Y. Zhang, five Chinese new record species, D. andersenii Lam,
D. sivanesanii C. Manoharachary et V. R. T. Reddy, E. frumentaeei (Mitra)
Leonard & Suggs, E. pedicellatium (Henry) Leonard &Suggs and eleven
species which had been reported in China, they are , D. avenacea
(M.A.Curtis ex Cooke) Shoem., O. dictyoides (Drechsler) Shoem.,
D. graminea (Rabenh. ex Schlecht.) Shoem., D. phlei (Graham) Shoem.,.
D. teres (Sate.) Shoem., E. fusiforme Alcom, E. holmii (Luttr.) Lenoard
& suggs, E. ]ongisporium G.Y.Sun, R.Zhang, M.Q.Zhu & T.Y.Zhang,
E. monoceras(Drechsler) Leonard & Suggs, E. phragmatis W.P Wu,
E. rostratum (Drechler) Leonard & Suggs and E. turcicum (Pass.) Leonard
& Suggs.
To investigate the genetic variation in E.rostratum, a total of 26
E.rostratum isolates and 7 isolates of other species were analyzed with 10
random amplified polymorphic DNA (RAPD) primers. The results showed
that there was a similar disorderliness with conidia morphology showed. There
was a large variation among all E.rostratum isolates, and character of conidia
was not a critical criteria of relationship between isolates. We should treat all
isolates of E.rostratum as one group since there were no constant crt. The conidia from culture medium shows less variety when
treated all isolates of E.rostratum as a whole, and should be treated as main
criteria of classification of this species.
After many works have been done about this two genera, the main results
of this research are the revisions of the Latin and Chinese name of the genera
and the species in them. We also have found a new species, several new
Chinese record species and broadened the hosts' records and distribution of
some species. All the works will contribute to the further study of this group of
fungi.
In our research work, we ve
payed a constant attention to the taxonomic
methods and the testifying and exploring of criteria to differentiate species. We
studied the culture character of all isolates, the conditions to spur Drechslera
spp to produce conidia on culture medium, the formation of conidia septum,
the germ character of conidia etc, and have concluded a much easier and
feasible criteria for the classification this group of fungi.
In differentiating different genera, the character of hilum plays an
important role. The hilum of Drechslera spp is atrium type hilum, which is
larger and we can see them never protruding from the contour of basal cell
when under optical microscope. Some time we can even see them were in the
basal cell of conidia, and clear of the cell wall. The hilum of Exserohilum spp.
were protruding and had an enveloping structure. But this structure can not be
found using optical microscope. Only that you may found an ring thickness at
the top of the hilum of E.monoceras and several other species. The
germination character of the basal cell of conidia is useful in distinguishing
Drechslera spp. from Bipolaris spp and Exserohilum spp But this character has
little difference between the later two genera. The color of conidia only has an
assitant role in genus classification. The Exserohilum spp have distoseptates
and the Curivularia spp with protruding hilum have euseptates has been seen
as the main difference between them. This criteria is used even it has been
doubted because there is no other criteria better.
In treating with the species, the morphology of conidia is main criteria,
especialy the shape and the size of conidia, the shape of conidia cell, the
conidia septum and the existence of secondary conidiophore and secondary
conidia. Some parameter such as the ratio of width of a
引文
1. 白逢彦. 真菌分子系统学研究简介(内部资料)[C]. 1999.
2. 白金铠. 长蠕孢菌 (Helminthosporium) 分类位置的变动[J]. 吉林农业大学学报, 1980,3: 13-19.
3. 蔡云鹏. 台湾植物病害名汇.(修订三版)[M]. 1991. 1～604页. 台湾:中华植物保护学会和植物病理学会出版.
4. 陈勇强等，国外微生物除草剂的研究及应用现状[J]，天津农业科学，1998, 4（2）：5-8。
5. 戴芳澜. 中国真菌总汇[M]. 1979. 科学出版社.
6. 邓叔群. 中国的真菌[M]. 1963. 1～808页.科学出版社
7. 董金皋等，玉米大斑病菌HT-毒素对玉米细胞VC氧化酶活性的影响[J]。河北农业大学学报，1999, 22（1）：50-53。
8. 冯洁，孙文姬，石磊岩，马存. 中国棉花枯萎菌生理小种的RAPD分析[J]. 菌物系统，2000，19（1）：45-50.
9. 葛起新.浙江植物病害志 病害篇(第一集)[M]. 1～328页及图版.上海科技出版社. 1991.
10. 姜广正.中国禾本科植物上的蠕形菌 (Helminthosporium) [J]. 植物病理学报. 5(1): 21-34, 1959.
11. 雷丽萍，白万明，李天飞，刘杏忠. RAPD在捕食性真菌遗传特性分析及分类鉴定中的应用[J]. 云南农业大学学报， 1998，13（1）：6-9.
12. 彭金火, 陆家云.内脐蠕孢属、平脐蠕孢属和突脐蠕孢属的研究[J]. 南京农业大学学报 , 1989, 12(4):46-52.
13. 戚佩坤等,吉林省栽培植物真菌病害志[M]. 1～479页. 科学出版社，1966.
14. 孙广宇.德氏霉属(Drechslera)及其相似属的分类研究[D]. 西北农业大学1988届攻读硕士学位研究生毕业论文.
15. 孙广宇等. 台湾省一些禾本科植物上蠕形菌的分类鉴定[J], 西北农业学报, 6(1): 92-93.
16. 王立秋，田新久.玉米大斑病研究现状及建议。黑龙江农业科学，1998, 4：32-33。
17. 王明旭译, 罗宽校. 长蠕孢属(Helminthosporium)种的分类[J]. 农业科技译丛, 1990. 19(1): 68-76.
18. 文景芝,陆家云. 内脐蠕孢属、平脐蠕孢属和突脐蠕孢属的分类鉴定[J]. 东北农学院学报, 1991. 22(2): 120-126.
19. 吴文平. 河北省丝孢菌的研究 Ⅰ 禾本科植物上蠕形菌的初步研究[J]. 河北省科学院学报, 1990, 1: 52～64
20. 阎培生，罗信昌，周启. 利用RAPDj技术对木耳属菌株进行分类鉴定的研究[J]. 菌物系统，2000， 19（1）：29-33.
21. 姚一建等，菌物学概论. 中国农业出版社, 2002.
22. 张天宇. 丝分孢子真菌(Mitosporic Fungi)的分类问题[C]. 山东农业大学学报(增刊) , 1999. 30:24-32.
23. 张修国等，烟草黑胫病菌株亲缘关系的RAPD分析[J]. 菌物系统， 2000， 19（1）：39-44.
24. 周永力，吕国忠，刘伟成，白金铠. 采用PCR-RAPD对球壳孢目真菌系统学的研究[J]. 菌物系统，1998, 17(2):160-166.
25. Abbas I.H.Resistance of barley stripe disease by mutation.Arab J.Plant protection[J], 1993,11(2):77-81.
26. Ainsworth, G.C. Ainsworth and Bisby's Dictionary of the Fungi[M]. CMI Kew. Surry. 1971.
27. Alcon J.L. Generic concepts in Drechslera,Bipolaris and Exserohilum[J]. Mycotaxon . 1983.Vol.XVII:1~86
28. Alcorn J.L. 1988. The taxonomy of Helminthosporium species. Ann.Rev.Phtopathol. 26:37~56.
29. Alcorn J.L. A new homothallice Setosphaerra species and its Exserohilum anamorph[J]. Trans.Br. mycol. Soc. 1985, 86(2): 313-317 .
30. Alcorn J.L. A new species of Exserohilum[J]. Trans.Br. mycol. Soc. 1988, 90(1):146-148.
31. Alcorn J.L. New combinations and synonymy in Bipolaris and Curvularia, and a new species of Exserohilum [J]. Mycotaxon, 1991, 41(2): 329-343.
32. Alcorn J.L. Setosphaeria monoceras sp.nov.,Ascigerous state of Exserohium monoceras[J]. Mycotaxon 1978, VII(2):411-414.
33. Alcorn J.L. Three Drechslera species on Chloris[J]. Trans. Br. mycol. Soc. 1976, 67:148-150.
34. Babadoost M. Sporulation of Drechslera graminea on barley straw extract agar[J]. Mycologia, 1998, 90(1):63-68.
35. Bakonyi J, Pomazi A, Hornok L. Molecular taxonomy of graminous species of the complex Helminthosporium[J]. Novenyvedelem, 1994, 30(5): 207-214.
36. Bar ZA, Tadmor Y, Shimoni M, Reuveni R. Resistance to norhern blight in maize(Zea mays) condtioned by the HtN gene and the assosiation with isoperoxidases[J]. Canadian Journal of Plant pathology,1998,20(1): 28-34.
37. Beth L.Teviotdale and Dennis H.Hall. Effect of light and temperature on number and length of Helminthosporium gramineum conidia produced in culture [J]. Can. J. Bot. 1976, 54: 644-648.
38. Borchardt D.S., Welz H.G. and Geiger H.H. Molecular marker analysis of European Setosphaeria turcica populations[J]. European Journal of Plant Pathology, 1998, 104: 611-617.
Borchardt DS, Welz HG, Geiger H. Molecular marker analysis of European
39. Setosphaeria tucica populations[J]. European Journal of Plant Pathology, 1998, 104: 611-617.
40. Braverman, S. W. Pigmentation and host range as species characters in Hilminthosporium.[J] Phytopathology 1957, 47:4
41. Braverman, S. W. The Helminthosporium gramineum complex and related species on cereals and forage grasses[J]. Phytopathology 1960, 50:688-695.
42. Carmichael J.W., Kendrick W.B., Connors I.L., Sigler L. Genera of Hyphomycetes. Edmonton: Univ. Alberta Press, 1980.
43. Castaneda R.R.E., Guarro J., Cano J. A new species of Exserohilum from Cuba[J]. Mycol.Res. 1995, 99(7): 825-826.
44. Chamberlain D.W., Allison J.L. The Brown leaf spot on Bromus inermis caused by Pyrenophora bromi[J]. Phytopathology 1945, 35:241-48
45. Chaudhuri S.. Validity of the 'type of conidial germination' as a taxonomic criterion in graminicolous Helminthosporium[J]. Bull. Indan Phytopathol. Soc. 1969, 5:15-19
46. Chidambaram P., Mathur S. B., Neergaard P. Identification of seedborn Drechslera species[J]. Friesia 1973, 10:165-207
47. Cooke D.E.L., Kennedy, D.M., Guy D.C., Russell J., Unkles S.G. and Duncan J.M. Relatedness of group I species of Phytophthora as assessed by randomly amplification polymorphic DNA (RAPD) and sequences of ribosomal DNA[J]. Mycol. Res. 1996, 100(3): 297-303.
48. Crous P.W., Janse B.J.H. and Tunbridge J. DNA homology between Pyrenophora japonica and P. teres [J]. Mycol. Res. 1995, 99(9): 1098-1102.
49. Danquah, O.A. Two new species of Drechslera[J]. Trans. Br. Mycol. Soc. 1975, 64:544-46
50. Dennis R.W.G. Studies in the morphology and biology of Helminthosporium avenae[J]. Trans. Br. Mycol. Soc. 1933, 18:223-38
51. Drechsler C. Phytopathological and taxonomic aspects of Ophiobolus, Pyrenophora, Helminthosporium, and a new genus, Cochliobolus[J]. Phytopathology 1934, 24: 953-983.
52. Drechsler, C. Some graminicolous species of Helminthosporium[J]: I. J. Agric. Res. 1923, 24:641-739
53. El Shafie A.E., Webster J. Setosphaeria khartoumensis sp. Nov. and its Exserohilum conidial state[J]. Trans. Br. Mycol. Soc. 1981, 77: 442.
54. Elliott E.S. 1949. The effect of the sugar concentration on conidial size of some species of Helminthosporium. Phytopathology 39: 953-58
55. Ellis M.B. Dematiacrous Hyphomycetes. C.M.I.，Kew. 1971.
56. Ellis M.B. More dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey, England :507, 1976.
57. El-Sayed AA. Disease resistance of barley genotypes under the conditions of northwestern coast of Egypt[J]. Assiut journal of agricultural sciences, 1991, 22(1):127-142.
58. Gatti.A. Physiological and biochemical variability in a population of Drechslera graminea[J].Journal of genitics and breeding, 1992,46(2):179-186.
59. Graham J.H. Helminthosporium leaf streak of timothy[J]. Phytopathology,1955, 45: 227-229
60. Hilu,HM. Host-pathogen Relationship of Helminthosporium turcicum in Resistant and Susceptible Corn seedlings[J].Pytopathology. 1964, 54:570-575.
61. Hiroshi.Biological control of rice leaf blast with Exserohilum monoceras, a pathogen of Echinochloa species[j],日本植物病理学会报，1999, 65（5）：543-548。
62. Honda Y. and Aragaki M. Stability of hilum protuberance in Exserohilum species[J]. Mycologia, 1978, 70: 547-555.
63. Hughes SJ. Conidiophores, conidia and classification [J]. Can. J. Bot., 1953, 31: 577-659.
64. Ito S. On some niw ascigerous stages of the species of Helminthosporium parsitic on cereals[J]. Proc. Imp. Acad. (Tokyo) 1930, 6:352-355
65. Janwhar M. Identification of Drechslera graminea isolates by cultural character and RAPD analysis[J].Cereal research communication. 2000, 28(1-2):87-93.
66. Jawhar M, Sangwan RS, Arabi MIE. Identification of Drechslera graminea isolates by cultural characters and RAPD analysis[J]. Cereal Research Communications,2000, 28:87-93.
67. Jooste, W. J. A new species of Drechslera on Tribulus terrestris[J]. Bothalia 1975, 11:511-13
68. Jorgensen HJL. Mechanisms of induced resistance in barley against Drechslera teres[J].Phytopahology, 1998, 88(7):698-707.
69. Kafi A. and Tarr A.J. Growth, sporulation and conidial characteristics of five graminicolous species of Helminthosporium [J]. Trans. Br. mycol. Soc. 1966, 49(2): 327-337.
70. Khetarpal R.K. A new species of Drechslera recorded on seed of Eleusine coracana [J]. Indian Phytopathology, 1984, 37: 320-321.
71. Lam A. Drechslera andersenii sp.nov. and other Drechslera sp. on ryegrass in England and Wales. Trans.Br.mycol.Soc. 1985, 85(4):595~602
72. Lam A. Drechslera siccans from ryegrass fields in England and wales. Trans. Br. mycol. Soc. 1984, 83(2): 305-311.
73. Lam A., Chapman P.F. Estimation of degree of similarity between species of Drechslera on Lolium spp. And Festuca pratensis using numerical taxonomy. Trans. Br. Mycol. Soc. 1985, 85: 603-614
74. Latterell F.M., Rossi A.E. 1973. Techniques for culture and photography that facilitate the identification of species of Helminthosporium. 2nd International Congress of Plant Pathology, Minneapolis: Abstracts of Papers No.153
75. Lawrie J, Greaves MP, Down VM. Drechslera sp. (Pyrenophora chaetomioides (Speg)), a potential biocontrol agent for Brimus sterilis and other Brumus spp[J]. Biocontrol Science and Technology, 1998,8(4):479-484.
76. Leach C.M., Fullerton R.A., Young K. Northern leaf blight of maize in New Zealand: relationship of Drechslera turcica airspora to factors influencing sporulation, conidium development, and chlamydospore formation. Phytopathology 1977, 67: 629-36
77. Leonard K.J. Setosphaeri prolata, the ascigerous state of Exserohilum prolatum [J]. Mycologia, 1974, 66: 281-297.
78. Leonard，KJ. Synonymy of Exserohilum halodes with E. rostratum，and induction of the ascigerous state，Setosphaeria rostrata. Mycologia 1976, 68:402-411.
79. Luttrell E.S. Correlations between conidial and acigerous state characters in Pyrenophora, Cochliobolus and Setosphaeria[J]. Revue de Mycologie, 1977, 41: 271-279.
80. Luttrell E.S. Taxonomic criteria in Helminthosporium [J]. Mycologia, 1963, 55: 643-674.
81. Luttrell ES. Approaches to the classification of Helminthosporium species[J]. Plant Dis. Rep. Suppl., 1954, 228: 111-113.
82. Luttrell, E. S. Leptosphaeria (Metasphaeria) perfect stages for Helminthosporium turcicum and H. rostratum. Phytopathology 1957, 47:313
83. Luttrell, E. S. Systematics of Helminthosporium and related genera. Mycologia ,1964, 56:119-32
84. Luttrell, E. SMorphology of Trichometasphaeria turcica. Am. J. Bot. . 1964, 51:213-19
85. Luttrell，E. S. A key to species of Helminthosporium reported on grasses in the United States. Pl. Dis. Rep. Suppl. 1951, 201:59-67.
86. Manoharachary C.,Thulasi Reddy V.R. A new species of Drechslera Ito from India. Current Science, 1983, 52(21):1026-1027.
87. McDonald, W. C. Variability and theinheritance of morphological mutants in Pyrenophora teres. Phytopathology 1967, 57:747-55
88. Michal Ond?ej. K vyskytu hub rod? Drechslera Ito a dendryphion wallroth(fugi imperfecti) na lnu v ?eskoslovensku. ?eská Mykologie, 1983, 37(2):72-76.
89. Michal Ond?ej. Key to the species Drechslera Ito sensu lato on the territory of Czechoslovakia. ?eská Mykologie, 1989, 43(1): 45-50.
90. Michal Ond?ej. Nové druhy hub rodu Drechslera Ito v ?eskoslovensku. ?eská Mykologie, 1988, 42(2): 84-89.
91. Misra, A.P. and Singh, R.A. Three new species of Drechslera from India [J]. Sydowia Annal. Mycol. Ser. 2, 1980, 32: 185-189.
92. Mitra, M., et al. A compatative study of species and strains of Helminthosporium on certain Indian cultivated crops [J]. Brit. Mycol. Soc. Trans., 1930, 15: 254-293.
93. Mostafa M.M. Biological control of Drechslera teres:ability of antagonists to reduce conidia formation ,coleoptile infection and leaf infection in barley(Hordeum vulgare),Mycolgie. 1993,14(4):287-295.
94. Mouchacca, J. Espèces nouvelles et communes de Drechslera, isolées de sols de regions arides [J]. Rev. Mycol., 1974, 38: 102-108.
95. Muchovej J.J.,Maria L.Ribeiro nesio. A new Exserohilum from Brazil[J]. Trans.Br. mycol. Soc. 1991, 89(1):126-128.
96. Muchovej, J J. Drechslera salviniae sp. nov. from Brazil [J]. Trans Br mycol Soc, 1979, 72(2): 331-332.
97. Mullin, B.A., C.G. Cook, and R.L. Schading. First report of Exserohilum rostratum, Curvularia senegalensis, and Nigrospora oryzae infecting kenaf in Texas. Pl. Dis. 1993, 77:846
98. Nelson. The perfect stage of Helminthosporium pedicellatum[J]. Mycologia, 1965, 57:665-668.
99. Nisikado, Y. Studies on the Helminthosporium diseases of Gramineae in Japan. Ber. Ohara Inst. Landwirtsch. Forsch. Kurashiki. 1929, 4:111-126 (English summary of 79)
100. Nolan.S.Control of Stagonospora nodorum and Septoria tritici in Wheat by pre-treatment with Drechslera teres, a non-host pathogen[J].European Journal of Plant Pathology, 2000, 106(2):203-207.
101. Odvody, G. N., Boosalis, M. G. An efficient, synchronous conidial production technique for Pyrenophora trichostoma. In Tan Spot of Wheat and Related Diseases Workshop, 1982. ed. R. M. Hosford, pp. 28-32. Fargo: North Dakota Agr. Exp. St.
102. Patil, A.S. and Rao, V.G. A new species of Drechslera from India[J], Trans. Br. mycol. Soc., 1972, 59(2): 339-340.
103. Paul A.R. Pyrenophora tetrarrhenae sp.nov.,Drechslera tetrarrhenae sp.nov.,and D.biseptata on Ehrharteae in Australia[J]. Trans.Br.mycol.Soc. 1971 ,56(2): 261-266.
104. Platt, H. W., Morrall, R. A. A., Gruen, H. E. The effects of substrate, temperature, and photoperiod on conidiation of Pyrenophora tritici-repentis. Can. J. Bot. 1977, 55:254-59
105. Reddy, S. M. Conidial germination in some isolates of Helminthosporium rostratum. Indian Phytopathol. 1975, 28:285-86
Richardson M.J., Fraser E.M. New combinations in Drechslera[J]. Trans Br mycol
106. Soc 1954, 51(1):147-148.
107. Ruiz Roldan.M.C. PTK1,a mitogen-activated-protein kinase gene,is required for conidiation,appressorium formation,and pathogenicity of Pyenosphora teres on barley[J].Molecular plant microbe interactions, 2001, 14(2):116-125.
108. Saccardo P A, Sylloge Fungorum. 1886-1972.
109. Samuel W B, Graham J H. Helminthosporium dictyoides and related species on forage grasses[J]. Phytopathology 1960, 50: 691~695
110. Schneider, D. J., Marasas, W. F. O., Collet, M. G., van der Westhuizen, G. C. A. An experimental mycotoxicosis in sheep and goats caused by Drechslera campanulata, a fungal pathogen of green oats[J]. Onderstepoort J. Vet. Res. 1985, 52:93-100
111. Shafie A.E.,Webster J. Setosphaeria khartoumensis sp.nov. and its Exserohilum conidial state. Trans Br mycol Soc, 1981, 77(2): 442~449
112. Shoemader, R. A. A pleomorphic parasite of cereal seeds, Pyrenophora semeniperda. Can. J. Bot. 1966, 44:1451-56
113. Shoemader, R. A. Drechslera Ito. Can. J. Bot. 1962, 40:809-36
114. Shoemaker, R. A. Marielliottia, a new genus of cereal and grass parasites segregated from Drechslera[J]. Can. J. Bot., 1998, 76: 1558-1569.
115. Shoemaker，R. A. Nomenclature of Drechslera and Bipolares，grass parasites degregated from 'Helminthosporium'[J]. Can. J. Bot. 1959, 37:879-887.
116. Sivanesan, A. Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs. CAB Internat. Mycol. Inst. Mycol. Pap. No.158. 1987.
117. Sivanesan, A. New Bipolaris, Curvularia and Exserohilum species[J]. Mycol. Res., 1992, 96(6): 485～489
118. Sivanesan, A. New Bipolaris, Curvularia and Exserohilum species[J]. Mycol. Res. 1992, 96:485-489
119. Sivanesan,A. New species of Exserohilum[J]. Trans. Br. Mycol. Soc. 1984 a, 83:319-329
120. Speakman, JB, Pommer, EH. A simple method for producing large volumes of Pyrenophora teres spore suspension[J]. Bull. Br. Mycol. Soc. 1986, 20:129-130
121. Spilker D.A.,Larsen P.O., et. Characterization and host range of Drechslera catenaria , the pathogen of leaf blight and crown rot of creeping bentgrass[J]. Plant Disease, 1985, 69(4):331-333.
122. Steiman R.,Giraud P. et. Exserohilum Israeli, a new species isolated from soil from Timna Park(Israel), and its physiological properties[J]. Antonie van Leeuwenhock. 2000, 78: 153-161.
123. Subramanian, CV, Jain B L. A revision of some graminicolous Helminthosporia[J]. Curr. Sci. 1966, 35: 352-355
124. Suggs Enda GSetosphaeria prolata, the ascigerous state of Exserohilum prolatum[J]. Mycologia, . 1974, 66 :281-297.
125. Sutton, B. C. Proposal for the conservation of the generic name Drechslera Ito (1930), Against Angiopoma Lev. (1841)[J]. Taxon 1977, 26: 591-592
126. Tautz，D. ( Rene，M. Simple sequences are ubiquitous repetitive components of eukaryotic genomes[J].Nucleic Acids. Res. 1984, 12: 4127-4138.
127. Teviotdale, BL, Hall, DH. Effect of light and temperature on number and length of Helminghosporium gramineum comidia produced in culture[J]. Can. J. Bot. 1976, 54:644-648
128.Upadhyay H.P. Dactylaria nervicola sp.nov.and Exserohilum novae-zelandiae comb.nob. form Mexico[J]. Mycologia, 1991, 83(3):371-376.
129.Wallwork H., Lichon A.,et. Drechslera wirreganensis-a new hyphomycete affecting barley in Australia[J]. Mycol.Res. 1992, 96(10):886-888.
130.Wallwork H., Lichon A.,et. Pyrenophora hordei—a new ascomycete with Drechslera anamorph affecting barley in Australia[J]. Mycol.Res., 1992, 96(12):1068-1070.
131.Welz H.G. Genes for resistance to northern corn leaf blight in diverse maize populations[J]. Plant Breeding, 2000,119:1-14.
132.Yuichi Honda. Stability of hilum protuberance in Exserohilum species[J]. Mycologia 1978, 70: 547-555.
133.Zhang WM. Host range of Exserohilum monoceras ,a potential bioherbicide for the control of Echinochloa species[J].Can.J.Bot. 1997, 75(5):685-692.
2. 白金铠. 长蠕孢菌 (Helminthosporium) 分类位置的变动[J]. 吉林农业大学学报, 1980,3: 13-19.
3. 蔡云鹏. 台湾植物病害名汇.(修订三版)[M]. 1991. 1～604页. 台湾:中华植物保护学会和植物病理学会出版.
4. 陈勇强等，国外微生物除草剂的研究及应用现状[J]，天津农业科学，1998, 4（2）：5-8。
5. 戴芳澜. 中国真菌总汇[M]. 1979. 科学出版社.
6. 邓叔群. 中国的真菌[M]. 1963. 1～808页.科学出版社
7. 董金皋等，玉米大斑病菌HT-毒素对玉米细胞VC氧化酶活性的影响[J]。河北农业大学学报，1999, 22（1）：50-53。
8. 冯洁，孙文姬，石磊岩，马存. 中国棉花枯萎菌生理小种的RAPD分析[J]. 菌物系统，2000，19（1）：45-50.
9. 葛起新.浙江植物病害志 病害篇(第一集)[M]. 1～328页及图版.上海科技出版社. 1991.
10. 姜广正.中国禾本科植物上的蠕形菌 (Helminthosporium) [J]. 植物病理学报. 5(1): 21-34, 1959.
11. 雷丽萍，白万明，李天飞，刘杏忠. RAPD在捕食性真菌遗传特性分析及分类鉴定中的应用[J]. 云南农业大学学报， 1998，13（1）：6-9.
12. 彭金火, 陆家云.内脐蠕孢属、平脐蠕孢属和突脐蠕孢属的研究[J]. 南京农业大学学报 , 1989, 12(4):46-52.
13. 戚佩坤等,吉林省栽培植物真菌病害志[M]. 1～479页. 科学出版社，1966.
14. 孙广宇.德氏霉属(Drechslera)及其相似属的分类研究[D]. 西北农业大学1988届攻读硕士学位研究生毕业论文.
15. 孙广宇等. 台湾省一些禾本科植物上蠕形菌的分类鉴定[J], 西北农业学报, 6(1): 92-93.
16. 王立秋，田新久.玉米大斑病研究现状及建议。黑龙江农业科学，1998, 4：32-33。
17. 王明旭译, 罗宽校. 长蠕孢属(Helminthosporium)种的分类[J]. 农业科技译丛, 1990. 19(1): 68-76.
18. 文景芝,陆家云. 内脐蠕孢属、平脐蠕孢属和突脐蠕孢属的分类鉴定[J]. 东北农学院学报, 1991. 22(2): 120-126.
19. 吴文平. 河北省丝孢菌的研究 Ⅰ 禾本科植物上蠕形菌的初步研究[J]. 河北省科学院学报, 1990, 1: 52～64
20. 阎培生，罗信昌，周启. 利用RAPDj技术对木耳属菌株进行分类鉴定的研究[J]. 菌物系统，2000， 19（1）：29-33.
21. 姚一建等，菌物学概论. 中国农业出版社, 2002.
22. 张天宇. 丝分孢子真菌(Mitosporic Fungi)的分类问题[C]. 山东农业大学学报(增刊) , 1999. 30:24-32.
23. 张修国等，烟草黑胫病菌株亲缘关系的RAPD分析[J]. 菌物系统， 2000， 19（1）：39-44.
24. 周永力，吕国忠，刘伟成，白金铠. 采用PCR-RAPD对球壳孢目真菌系统学的研究[J]. 菌物系统，1998, 17(2):160-166.
25. Abbas I.H.Resistance of barley stripe disease by mutation.Arab J.Plant protection[J], 1993,11(2):77-81.
26. Ainsworth, G.C. Ainsworth and Bisby's Dictionary of the Fungi[M]. CMI Kew. Surry. 1971.
27. Alcon J.L. Generic concepts in Drechslera,Bipolaris and Exserohilum[J]. Mycotaxon . 1983.Vol.XVII:1~86
28. Alcorn J.L. 1988. The taxonomy of Helminthosporium species. Ann.Rev.Phtopathol. 26:37~56.
29. Alcorn J.L. A new homothallice Setosphaerra species and its Exserohilum anamorph[J]. Trans.Br. mycol. Soc. 1985, 86(2): 313-317 .
30. Alcorn J.L. A new species of Exserohilum[J]. Trans.Br. mycol. Soc. 1988, 90(1):146-148.
31. Alcorn J.L. New combinations and synonymy in Bipolaris and Curvularia, and a new species of Exserohilum [J]. Mycotaxon, 1991, 41(2): 329-343.
32. Alcorn J.L. Setosphaeria monoceras sp.nov.,Ascigerous state of Exserohium monoceras[J]. Mycotaxon 1978, VII(2):411-414.
33. Alcorn J.L. Three Drechslera species on Chloris[J]. Trans. Br. mycol. Soc. 1976, 67:148-150.
34. Babadoost M. Sporulation of Drechslera graminea on barley straw extract agar[J]. Mycologia, 1998, 90(1):63-68.
35. Bakonyi J, Pomazi A, Hornok L. Molecular taxonomy of graminous species of the complex Helminthosporium[J]. Novenyvedelem, 1994, 30(5): 207-214.
36. Bar ZA, Tadmor Y, Shimoni M, Reuveni R. Resistance to norhern blight in maize(Zea mays) condtioned by the HtN gene and the assosiation with isoperoxidases[J]. Canadian Journal of Plant pathology,1998,20(1): 28-34.
37. Beth L.Teviotdale and Dennis H.Hall. Effect of light and temperature on number and length of Helminthosporium gramineum conidia produced in culture [J]. Can. J. Bot. 1976, 54: 644-648.
38. Borchardt D.S., Welz H.G. and Geiger H.H. Molecular marker analysis of European Setosphaeria turcica populations[J]. European Journal of Plant Pathology, 1998, 104: 611-617.
Borchardt DS, Welz HG, Geiger H. Molecular marker analysis of European
39. Setosphaeria tucica populations[J]. European Journal of Plant Pathology, 1998, 104: 611-617.
40. Braverman, S. W. Pigmentation and host range as species characters in Hilminthosporium.[J] Phytopathology 1957, 47:4
41. Braverman, S. W. The Helminthosporium gramineum complex and related species on cereals and forage grasses[J]. Phytopathology 1960, 50:688-695.
42. Carmichael J.W., Kendrick W.B., Connors I.L., Sigler L. Genera of Hyphomycetes. Edmonton: Univ. Alberta Press, 1980.
43. Castaneda R.R.E., Guarro J., Cano J. A new species of Exserohilum from Cuba[J]. Mycol.Res. 1995, 99(7): 825-826.
44. Chamberlain D.W., Allison J.L. The Brown leaf spot on Bromus inermis caused by Pyrenophora bromi[J]. Phytopathology 1945, 35:241-48
45. Chaudhuri S.. Validity of the 'type of conidial germination' as a taxonomic criterion in graminicolous Helminthosporium[J]. Bull. Indan Phytopathol. Soc. 1969, 5:15-19
46. Chidambaram P., Mathur S. B., Neergaard P. Identification of seedborn Drechslera species[J]. Friesia 1973, 10:165-207
47. Cooke D.E.L., Kennedy, D.M., Guy D.C., Russell J., Unkles S.G. and Duncan J.M. Relatedness of group I species of Phytophthora as assessed by randomly amplification polymorphic DNA (RAPD) and sequences of ribosomal DNA[J]. Mycol. Res. 1996, 100(3): 297-303.
48. Crous P.W., Janse B.J.H. and Tunbridge J. DNA homology between Pyrenophora japonica and P. teres [J]. Mycol. Res. 1995, 99(9): 1098-1102.
49. Danquah, O.A. Two new species of Drechslera[J]. Trans. Br. Mycol. Soc. 1975, 64:544-46
50. Dennis R.W.G. Studies in the morphology and biology of Helminthosporium avenae[J]. Trans. Br. Mycol. Soc. 1933, 18:223-38
51. Drechsler C. Phytopathological and taxonomic aspects of Ophiobolus, Pyrenophora, Helminthosporium, and a new genus, Cochliobolus[J]. Phytopathology 1934, 24: 953-983.
52. Drechsler, C. Some graminicolous species of Helminthosporium[J]: I. J. Agric. Res. 1923, 24:641-739
53. El Shafie A.E., Webster J. Setosphaeria khartoumensis sp. Nov. and its Exserohilum conidial state[J]. Trans. Br. Mycol. Soc. 1981, 77: 442.
54. Elliott E.S. 1949. The effect of the sugar concentration on conidial size of some species of Helminthosporium. Phytopathology 39: 953-58
55. Ellis M.B. Dematiacrous Hyphomycetes. C.M.I.，Kew. 1971.
56. Ellis M.B. More dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey, England :507, 1976.
57. El-Sayed AA. Disease resistance of barley genotypes under the conditions of northwestern coast of Egypt[J]. Assiut journal of agricultural sciences, 1991, 22(1):127-142.
58. Gatti.A. Physiological and biochemical variability in a population of Drechslera graminea[J].Journal of genitics and breeding, 1992,46(2):179-186.
59. Graham J.H. Helminthosporium leaf streak of timothy[J]. Phytopathology,1955, 45: 227-229
60. Hilu,HM. Host-pathogen Relationship of Helminthosporium turcicum in Resistant and Susceptible Corn seedlings[J].Pytopathology. 1964, 54:570-575.
61. Hiroshi.Biological control of rice leaf blast with Exserohilum monoceras, a pathogen of Echinochloa species[j],日本植物病理学会报，1999, 65（5）：543-548。
62. Honda Y. and Aragaki M. Stability of hilum protuberance in Exserohilum species[J]. Mycologia, 1978, 70: 547-555.
63. Hughes SJ. Conidiophores, conidia and classification [J]. Can. J. Bot., 1953, 31: 577-659.
64. Ito S. On some niw ascigerous stages of the species of Helminthosporium parsitic on cereals[J]. Proc. Imp. Acad. (Tokyo) 1930, 6:352-355
65. Janwhar M. Identification of Drechslera graminea isolates by cultural character and RAPD analysis[J].Cereal research communication. 2000, 28(1-2):87-93.
66. Jawhar M, Sangwan RS, Arabi MIE. Identification of Drechslera graminea isolates by cultural characters and RAPD analysis[J]. Cereal Research Communications,2000, 28:87-93.
67. Jooste, W. J. A new species of Drechslera on Tribulus terrestris[J]. Bothalia 1975, 11:511-13
68. Jorgensen HJL. Mechanisms of induced resistance in barley against Drechslera teres[J].Phytopahology, 1998, 88(7):698-707.
69. Kafi A. and Tarr A.J. Growth, sporulation and conidial characteristics of five graminicolous species of Helminthosporium [J]. Trans. Br. mycol. Soc. 1966, 49(2): 327-337.
70. Khetarpal R.K. A new species of Drechslera recorded on seed of Eleusine coracana [J]. Indian Phytopathology, 1984, 37: 320-321.
71. Lam A. Drechslera andersenii sp.nov. and other Drechslera sp. on ryegrass in England and Wales. Trans.Br.mycol.Soc. 1985, 85(4):595~602
72. Lam A. Drechslera siccans from ryegrass fields in England and wales. Trans. Br. mycol. Soc. 1984, 83(2): 305-311.
73. Lam A., Chapman P.F. Estimation of degree of similarity between species of Drechslera on Lolium spp. And Festuca pratensis using numerical taxonomy. Trans. Br. Mycol. Soc. 1985, 85: 603-614
74. Latterell F.M., Rossi A.E. 1973. Techniques for culture and photography that facilitate the identification of species of Helminthosporium. 2nd International Congress of Plant Pathology, Minneapolis: Abstracts of Papers No.153
75. Lawrie J, Greaves MP, Down VM. Drechslera sp. (Pyrenophora chaetomioides (Speg)), a potential biocontrol agent for Brimus sterilis and other Brumus spp[J]. Biocontrol Science and Technology, 1998,8(4):479-484.
76. Leach C.M., Fullerton R.A., Young K. Northern leaf blight of maize in New Zealand: relationship of Drechslera turcica airspora to factors influencing sporulation, conidium development, and chlamydospore formation. Phytopathology 1977, 67: 629-36
77. Leonard K.J. Setosphaeri prolata, the ascigerous state of Exserohilum prolatum [J]. Mycologia, 1974, 66: 281-297.
78. Leonard，KJ. Synonymy of Exserohilum halodes with E. rostratum，and induction of the ascigerous state，Setosphaeria rostrata. Mycologia 1976, 68:402-411.
79. Luttrell E.S. Correlations between conidial and acigerous state characters in Pyrenophora, Cochliobolus and Setosphaeria[J]. Revue de Mycologie, 1977, 41: 271-279.
80. Luttrell E.S. Taxonomic criteria in Helminthosporium [J]. Mycologia, 1963, 55: 643-674.
81. Luttrell ES. Approaches to the classification of Helminthosporium species[J]. Plant Dis. Rep. Suppl., 1954, 228: 111-113.
82. Luttrell, E. S. Leptosphaeria (Metasphaeria) perfect stages for Helminthosporium turcicum and H. rostratum. Phytopathology 1957, 47:313
83. Luttrell, E. S. Systematics of Helminthosporium and related genera. Mycologia ,1964, 56:119-32
84. Luttrell, E. SMorphology of Trichometasphaeria turcica. Am. J. Bot. . 1964, 51:213-19
85. Luttrell，E. S. A key to species of Helminthosporium reported on grasses in the United States. Pl. Dis. Rep. Suppl. 1951, 201:59-67.
86. Manoharachary C.,Thulasi Reddy V.R. A new species of Drechslera Ito from India. Current Science, 1983, 52(21):1026-1027.
87. McDonald, W. C. Variability and theinheritance of morphological mutants in Pyrenophora teres. Phytopathology 1967, 57:747-55
88. Michal Ond?ej. K vyskytu hub rod? Drechslera Ito a dendryphion wallroth(fugi imperfecti) na lnu v ?eskoslovensku. ?eská Mykologie, 1983, 37(2):72-76.
89. Michal Ond?ej. Key to the species Drechslera Ito sensu lato on the territory of Czechoslovakia. ?eská Mykologie, 1989, 43(1): 45-50.
90. Michal Ond?ej. Nové druhy hub rodu Drechslera Ito v ?eskoslovensku. ?eská Mykologie, 1988, 42(2): 84-89.
91. Misra, A.P. and Singh, R.A. Three new species of Drechslera from India [J]. Sydowia Annal. Mycol. Ser. 2, 1980, 32: 185-189.
92. Mitra, M., et al. A compatative study of species and strains of Helminthosporium on certain Indian cultivated crops [J]. Brit. Mycol. Soc. Trans., 1930, 15: 254-293.
93. Mostafa M.M. Biological control of Drechslera teres:ability of antagonists to reduce conidia formation ,coleoptile infection and leaf infection in barley(Hordeum vulgare),Mycolgie. 1993,14(4):287-295.
94. Mouchacca, J. Espèces nouvelles et communes de Drechslera, isolées de sols de regions arides [J]. Rev. Mycol., 1974, 38: 102-108.
95. Muchovej J.J.,Maria L.Ribeiro nesio. A new Exserohilum from Brazil[J]. Trans.Br. mycol. Soc. 1991, 89(1):126-128.
96. Muchovej, J J. Drechslera salviniae sp. nov. from Brazil [J]. Trans Br mycol Soc, 1979, 72(2): 331-332.
97. Mullin, B.A., C.G. Cook, and R.L. Schading. First report of Exserohilum rostratum, Curvularia senegalensis, and Nigrospora oryzae infecting kenaf in Texas. Pl. Dis. 1993, 77:846
98. Nelson. The perfect stage of Helminthosporium pedicellatum[J]. Mycologia, 1965, 57:665-668.
99. Nisikado, Y. Studies on the Helminthosporium diseases of Gramineae in Japan. Ber. Ohara Inst. Landwirtsch. Forsch. Kurashiki. 1929, 4:111-126 (English summary of 79)
100. Nolan.S.Control of Stagonospora nodorum and Septoria tritici in Wheat by pre-treatment with Drechslera teres, a non-host pathogen[J].European Journal of Plant Pathology, 2000, 106(2):203-207.
101. Odvody, G. N., Boosalis, M. G. An efficient, synchronous conidial production technique for Pyrenophora trichostoma. In Tan Spot of Wheat and Related Diseases Workshop, 1982. ed. R. M. Hosford, pp. 28-32. Fargo: North Dakota Agr. Exp. St.
102. Patil, A.S. and Rao, V.G. A new species of Drechslera from India[J], Trans. Br. mycol. Soc., 1972, 59(2): 339-340.
103. Paul A.R. Pyrenophora tetrarrhenae sp.nov.,Drechslera tetrarrhenae sp.nov.,and D.biseptata on Ehrharteae in Australia[J]. Trans.Br.mycol.Soc. 1971 ,56(2): 261-266.
104. Platt, H. W., Morrall, R. A. A., Gruen, H. E. The effects of substrate, temperature, and photoperiod on conidiation of Pyrenophora tritici-repentis. Can. J. Bot. 1977, 55:254-59
105. Reddy, S. M. Conidial germination in some isolates of Helminthosporium rostratum. Indian Phytopathol. 1975, 28:285-86
Richardson M.J., Fraser E.M. New combinations in Drechslera[J]. Trans Br mycol
106. Soc 1954, 51(1):147-148.
107. Ruiz Roldan.M.C. PTK1,a mitogen-activated-protein kinase gene,is required for conidiation,appressorium formation,and pathogenicity of Pyenosphora teres on barley[J].Molecular plant microbe interactions, 2001, 14(2):116-125.
108. Saccardo P A, Sylloge Fungorum. 1886-1972.
109. Samuel W B, Graham J H. Helminthosporium dictyoides and related species on forage grasses[J]. Phytopathology 1960, 50: 691~695
110. Schneider, D. J., Marasas, W. F. O., Collet, M. G., van der Westhuizen, G. C. A. An experimental mycotoxicosis in sheep and goats caused by Drechslera campanulata, a fungal pathogen of green oats[J]. Onderstepoort J. Vet. Res. 1985, 52:93-100
111. Shafie A.E.,Webster J. Setosphaeria khartoumensis sp.nov. and its Exserohilum conidial state. Trans Br mycol Soc, 1981, 77(2): 442~449
112. Shoemader, R. A. A pleomorphic parasite of cereal seeds, Pyrenophora semeniperda. Can. J. Bot. 1966, 44:1451-56
113. Shoemader, R. A. Drechslera Ito. Can. J. Bot. 1962, 40:809-36
114. Shoemaker, R. A. Marielliottia, a new genus of cereal and grass parasites segregated from Drechslera[J]. Can. J. Bot., 1998, 76: 1558-1569.
115. Shoemaker，R. A. Nomenclature of Drechslera and Bipolares，grass parasites degregated from 'Helminthosporium'[J]. Can. J. Bot. 1959, 37:879-887.
116. Sivanesan, A. Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs. CAB Internat. Mycol. Inst. Mycol. Pap. No.158. 1987.
117. Sivanesan, A. New Bipolaris, Curvularia and Exserohilum species[J]. Mycol. Res., 1992, 96(6): 485～489
118. Sivanesan, A. New Bipolaris, Curvularia and Exserohilum species[J]. Mycol. Res. 1992, 96:485-489
119. Sivanesan,A. New species of Exserohilum[J]. Trans. Br. Mycol. Soc. 1984 a, 83:319-329
120. Speakman, JB, Pommer, EH. A simple method for producing large volumes of Pyrenophora teres spore suspension[J]. Bull. Br. Mycol. Soc. 1986, 20:129-130
121. Spilker D.A.,Larsen P.O., et. Characterization and host range of Drechslera catenaria , the pathogen of leaf blight and crown rot of creeping bentgrass[J]. Plant Disease, 1985, 69(4):331-333.
122. Steiman R.,Giraud P. et. Exserohilum Israeli, a new species isolated from soil from Timna Park(Israel), and its physiological properties[J]. Antonie van Leeuwenhock. 2000, 78: 153-161.
123. Subramanian, CV, Jain B L. A revision of some graminicolous Helminthosporia[J]. Curr. Sci. 1966, 35: 352-355
124. Suggs Enda GSetosphaeria prolata, the ascigerous state of Exserohilum prolatum[J]. Mycologia, . 1974, 66 :281-297.
125. Sutton, B. C. Proposal for the conservation of the generic name Drechslera Ito (1930), Against Angiopoma Lev. (1841)[J]. Taxon 1977, 26: 591-592
126. Tautz，D. ( Rene，M. Simple sequences are ubiquitous repetitive components of eukaryotic genomes[J].Nucleic Acids. Res. 1984, 12: 4127-4138.
127. Teviotdale, BL, Hall, DH. Effect of light and temperature on number and length of Helminghosporium gramineum comidia produced in culture[J]. Can. J. Bot. 1976, 54:644-648
128.Upadhyay H.P. Dactylaria nervicola sp.nov.and Exserohilum novae-zelandiae comb.nob. form Mexico[J]. Mycologia, 1991, 83(3):371-376.
129.Wallwork H., Lichon A.,et. Drechslera wirreganensis-a new hyphomycete affecting barley in Australia[J]. Mycol.Res. 1992, 96(10):886-888.
130.Wallwork H., Lichon A.,et. Pyrenophora hordei—a new ascomycete with Drechslera anamorph affecting barley in Australia[J]. Mycol.Res., 1992, 96(12):1068-1070.
131.Welz H.G. Genes for resistance to northern corn leaf blight in diverse maize populations[J]. Plant Breeding, 2000,119:1-14.
132.Yuichi Honda. Stability of hilum protuberance in Exserohilum species[J]. Mycologia 1978, 70: 547-555.
133.Zhang WM. Host range of Exserohilum monoceras ,a potential bioherbicide for the control of Echinochloa species[J].Can.J.Bot. 1997, 75(5):685-692.